Primer

The development of
motor behavior
Karen E. Adolph1* and John M. Franchak2

This article reviews research on the development of motor behavior from a

developmental systems perspective. We focus on infancy when basic action sys-
tems are acquired. Posture provides a stable base for locomotion, manual
actions, and facial actions. Experience facilitates improvements in motor behav-
ior and infants accumulate immense amounts of experience with all of their basic
action systems. At every point in development, perception guides motor behav-
ior by providing feedback about the results of just prior movements and infor-
mation about what to do next. Reciprocally, the development of motor behavior
provides fodder for perception. More generally, motor development brings about
new opportunities for acquiring knowledge about the world, and burgeoning
motor skills can instigate cascades of developmental changes in perceptual, cog-
nitive, and social domains. © 2016 Wiley Periodicals, Inc.
How to cite this article:
WIREs Cogn Sci 2016. doi: 10.1002/wcs.1430

INTRODUCTION Movements generate perceptual information, provide

the means for acquiring knowledge about the world,

M otor behavior includes every kind of movement

from involuntary twitches to goal-directed
actions, in every part of the body from head to toe,
in every physical and social context from solitary
play to group interactions. The development of
motor behavior bridges the entire life span from the
first fetal movement to the last dying breath.
Although movements fundamentally depend on
generating, controlling, and exploiting physical
forces, managing forces requires more than muscles
and biomechanics. At every point in development,
adaptive control of movement relies on core psycho-
logical functions.1,2 Perception and cognition are
required to plan and guide actions.3 Social and cul-
tural factors spur and constrain motor behaviors.4
Motor behaviors, in turn, provide the raw material
for perception, cognition, and social interaction.5,6
*Correspondence to: karen.adolph@nyu.edu
1 with their children affect the form of new skills, the
Department of Psychology, New York University, New York,
NY, USA
2
Department of Psychology, University of California, Riverside,
CA, USA
Conflict of interest: The authors have declared no conflicts of inter-
est for this article.
and make social interactions possible.
According to a developmental systems view (see
Blumberg, Development evolving: the origins and
meanings of instinct, and Lickliter, Developmental
evolution, WIREs Cogn Sci, also in the collection
How We Develop), motor behaviors cannot be
understood in isolation, divorced from the bodily,
environmental, and social/cultural context in which
they occur.7 Movements are inextricably nested in a
body-environment system (see Oudeyer, What do we
learn about development from baby robots?, WIREs
Cogn Sci, also in the collection How We Develop).
The body and the environment develop in tandem.
New or improved motor skills bring new parts of the
environment into play and thereby provide new or
enhanced opportunities for learning and doing. Care-
giving practices facilitate and constrain motor devel-
opment. As a consequence, differences in the way
caregivers structure the environment and interact
ages when they first appear, and the shape of their
developmental trajectories.
New motor behaviors can emerge from a mix
of interacting factors, some so pervasive that we mis-
takenly take them for granted, and some so subtle or

© 2016 Wiley Periodicals, Inc.

 Primer
nonobvious that we fail to recognize the link. Devel-
opmental changes in one domain can have cascading
effects on development in other domains, sometimes
far afield from the original accomplishment.8,9 More-
over, the context in which behavior develops can be
very different for individual children, resulting in
developmental pathways that sometimes converge at
the same outcome and sometimes veer off in unique
directions.
This article is organized around four basic
action systems—posture, locomotion, manual
actions, and movements in the face and head. We
focus primarily on the infancy period, when basic
action systems are acquired.
POSTURE
Posture is the most fundamental of motor actions. It
is the foundation upon which other actions are
built.10 The instant that any part of the body breaks
from the support surface—merely raising an arm
while supine or lifting the head while prone—torque
acting on the body part creates disequilibrium. This
is why novice sitting and standing infants lose bal-
ance just from turning their heads or lifting their
arms. Posture must be sufficiently stable to allow
movements of the extremities, and maintaining a sta-
ble posture sets up the necessary conditions for look-
ing around, handling objects, holding conversations,
or going somewhere. As such, the emergence of most
skills—including those not obviously related to
posture—must await the development of sufficient
postural control. Like every action, posture is percep-
tually guided and maintained.
Overcoming Gravity
Gravity and the surrounding media (e.g., air, water,
the ground beneath the feet) are so quietly pervasive,
so hidden in plain sight, that these important factors
are often overlooked as causal forces in development.
But they are central for motor development. Before
birth, the buoyant uterine environment supports a
variety of postures. Large body movements—whole
body flexion and extension, stretching and writhing,
and vigorous leg kicks that somersault the fetus
through the amniotic fluid—peak at 14–16 weeks
gestation.11,12 As the growing fetus occupies increas-
ingly more space in the uterus, the propensity for
movement is masked until the fetus can no longer
extend its limbs or turn its head. Many of the move-
ments practiced by the fetus are present in the reper-
toire of the neonate,12 but after birth begins the real
struggle against gravity.
© 2016 Wiley Periodicals, Inc.
wires.wiley.com/cogsci
Postural development is the attainment of
increasingly erect postures poised over an increasingly
small base of support. Think of a newborn struggling
to lift its head, a toddler’s wide walking stance, and an
older child dancing on pointe. Indeed, the most com-
mon images of motor development are milestone
charts of postural development (Figure 1). The mile-
stone charts suggest an orderly, age-related march
through a series of stages, but developmental pathways
can differ and individual infants do not strictly adhere
to the normative sequence derived from average onset
ages. Infants can acquire skills in various orders, skip
stages, and revert to earlier forms.13–15 Moreover, the
skills highlighted on the milestone charts reflect the cul-
tural biases of the initial researchers and samples.4 In
some cultures, for example, many infants do not crawl,
or they do so after they learn to walk.16
Generally speaking, infants’ gradual triumph
over gravity precedes top down from head to feet.
The top-down progression is especially striking in the
development of sitting. At first, head and trunk con-
trol is so poor that unsupported infants fold in half,
falling chest to legs. Increasing control moves slowly
down the spine—neck, shoulders, waist, and hips.17,18
Infants eventually ‘tripod sit’ by stabilizing their torso
with arms propped between their outstretched legs.
Finally, around 6 months of age, infants sit independ-
ently with hands freed from a supporting role,19 and
over the ensuing weeks gain sufficient stability to
manage the destabilizing forces caused by turning the
head, twisting the torso, and moving the arms. The
skill progression is not locked to a strict maturational
timetable. Differences in childrearing practices affect
the timing and trajectory of sitting.4,20 In cultures
where caregivers routinely exercise and massage their
infants, the babies sit independently before 5 months
of age, and they do so with such assured stability that
their mothers regularly perch them on high furniture
and leave the room to do chores.
Like sitting, standing typically begins with man-
ual support of balance. Infants pull to stand and hold
themselves upright by gripping furniture for sup-
port.15 Toward the end of the first year, they stand
freely and cruise holding furniture for support.19
Locomotion in prone, sitting, crawling, and upright
postures appears only after infants can keep balance
in one place, and transitions between postures (shift-
ing from prone to sitting, sitting to standing, and so
on) typically emerge last.
Basis for Action
A stable postural base opens up new possibilities for
acquiring knowledge and acting on the world. The
 WIREs Cognitive Science The development of motor behavior

Walks alone

Stands alone

Cruising

Pulls to stand

Stands with support

Crawling

Sits without support

Rolls over

Prone, Chest up; Uses arms for support

Prone, Lifts head

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16
Age (months)

FI GU RE 1 | Typical example of milestone chart illustrating age-related changes in postural development. (Reprinted with permission from Ref
4. Copyright 2010 Taylor and Francis)

ability to maintain head position while held in care-

givers’ arms allows infants to look around21 and
maintain gaze with caregivers.22 The ability to sit
and stand upright provides new vantage points for
visual exploration.23 Stability in a sitting posture
frees the arms for reaching and the hands for manual
exploration.17,24–26
Indeed, reaching and manual exploration have
different developmental trajectories for prone, supine,
and sitting postures.25,27 While prone, bimanual
exploration is difficult because one arm is occupied
in holding the chest off the floor. While supine, F I G U R E 2 | Toddler losing balance in a ‘moving room.’ Child
infants struggle to raise objects against gravity and stands on a solid floor surrounded by walls that move back and forth
have difficulty exploring them visually. But while sit- along a track. Here, the walls move toward the child creating the
ting, more sophisticated bimanual object exploration visual illusion of the body swaying forward; the child compensates by
swaying backward. (Reprinted with permission from Ref 37. Copyright
is possible because head, arms, and hands are more
1989 American Psychological Association)
free to move.
Developmental changes in postural control
instigate a cascade of far-flung changes: Independent Dynamic Postural Control
sitting facilitates more sophisticated bimanual object Movement is ubiquitous in every posture. Even while
exploration such as fingering, transferring, and lying down, the body is in motion. Similarly, sitting,
rotating, which in turn facilitate learning about the crawling, and standing postures may appear station-
three-dimensionality of objects.28 Improvements in ary to casual observation, but they are not. Rather,
manual skills are also linked with shifts in infants’ the body gently sways back and forth within the base
attention to changes in object appearance,29 object of support.18,34 A torque-induced sway in one direc-
size,30 multimodal information about objects,31 and tion must be met by a muscle-induced compensatory
other people’s intentions to grasp objects.32,33 The sway in the opposite direction. Standing infants are
path from posture to prehension to perceptual learn- sensitive to perceptual information for body sway
ing is not immediately obvious, but it is there and can control swaying movements with merely a
nonetheless. light touch of the hand on a support surface.35,36

© 2016 Wiley Periodicals, Inc.

 Primer
Visual information for body sway is extremely
powerful. Slight movement of the walls around a sit-
ting or standing infant in a ‘moving room’ creates the
illusion of postural sway (Figure 2), and infants com-
pensate for the visual information for body position
by leaning in the opposite direction.37–39 However,
infants do not use visual information for postural con-
trol as efficiently as older children and adults. While
adults rock gently back and forth like puppets in tune
with the wall oscillations, infants’ compensatory
sways are excessive and they often stagger and fall.
Summary: Posture
Posture is the core ingredient of motor skill. With no
postural control, most motor behaviors are impossi-
ble. The development of postural control instigates a
cascade of new skills and opens up new possibilities
for looking, social interactions, manual actions, and
locomotion. Postural development is partly a percep-
tual accomplishment because even while sitting and
standing, the body is always slightly swaying and
perception plays a key role in keeping the body inside
the base of support. Postural control emerges from
the interaction of a growing body dealing with the
constraints of the physical environment—gravity, air,
the properties of the support surface, and so
on. Caregiving practices can speed up or delay pos-
tural control and the cascade of new skills that
follow.
LOCOMOTION
Precursory locomotor movements are exhibited dur-
ing fetal and neonatal periods, but locomotion is not
reflexive or hardwired. Rather, locomotion is creative
and infants must learn to control locomotion adapt-
ively. Locomotion improves with practice, and prac-
tice can lead to extraordinary performance.4,7,40
Newborn Reflexes
When newborns are held upright with their feet on a
hard surface, they move their legs in an alternating
pattern that resembles walking. This phenomenon is
called the ‘newborn stepping reflex’ because the
movements appear to be elicited by contact with the
ground surface and do not require cortical control.41
Stepping typically disappears by 2 months of age and
reappears at 8–10 months when infants begin walk-
ing with support. The fact that newborns produce
alternating, upright leg movements led researchers to
believe that walking is hardwired in the nervous
system.42–44 Similarly, the curious disappearance and
© 2016 Wiley Periodicals, Inc.
wires.wiley.com/cogsci
reappearance of stepping was attributed to a hard-
wired developmental mechanism: Cortical matura-
tion inhibits the reflex and increased myelination of
the corticospinal tract allows stepping to return
under volitional control.
However, the so-called stepping reflex is not, in
fact, reflexive, and alternating leg movements do not,
in fact, disappear. Newborns ‘air-step’ without an eli-
citing physical stimulus and they step in response to
optic flow.45,46 Infants can deliberately modify their
leg movements47 in various configurations of alter-
nating, simultaneous, and single-leg kicks.48,49 They
spontaneously kick their legs while supine50 and
supine leg kicks are kinematically equivalent to
upright steps, and are produced by the same muscle
activations.50 Moreover, supine kicking continues
unabated throughout the period when upright step-
ping disappears,51 and upright steps instantly reap-
pear when infants are held on a motorized
treadmill52,53 or when their legs are submerged in a
tank of water.54 With daily practice in an upright
posture, the stepping movements never disap-
pear.55,56 Changes in the body, not the brain, explain
the U-shaped trajectory of upright stepping: Between
2–8 months of age, gains in leg fat typically outstrip
gains in muscle.54 In an upright position, infants can-
not lift their chubby legs against gravity, but in a
supine position gravity helps to flex the legs; on a
treadmill, the moving belt does the work of pulling
infants’ legs backward and in a tank of water, the
medium alleviates the effects of gravity. Upright prac-
tice makes leg muscles stronger.
Creative Solutions
Individual infants find different ways to solve the
problem of moving. Their first success at mobility
likely involves a prone position with minimal balance
constraints. They may log roll from place to place or
pivot in circles using auditory information to calcu-
late the shortest rotational distance to their care-
givers.57 As shown in Figure 3, some infants belly
crawl, using limbs, head, and belly in various combi-
nations for support and propulsion.58,59 The belly
rests continually on the floor or bumps up and down
during each cycle. Every form of precursory prone
movement helps: Infants who pivot, belly crawl, and
so on are twice as proficient when they begin crawl-
ing on hands and knees compared with infants who
do not display the earlier forms.58 In fact, simply
spending a few minutes a day in a prone position
accelerates the onset of rolling and crawling.60
On hands and knees, balance constraints
increase because the belly is off the floor. As a
 WIREs Cognitive Science The development of motor behavior

(a)

Crawl Sit Prone Backing

(b)

Hunchback Windsurfing Mountain-climbing Drunken

FI G URE 4 | (A) Some of the strategies infants use to descend

slopes: Scooting down in a sitting position, crawling on hands and
knees, sliding head-first while prone, and turning their bodies to back
down feet first. (B) Some of the strategies infants use to cross bridges
holding a wobbly handrail for support. Infants employ a ‘hunchback’
strategy by pushing down on the rail to make it taut, walk sideways
while leaning backward as if ‘windsurfing,’ walking forward and
pulling back on the rail as if ‘mountain climbing,’ and ‘drunkenly’
leaning against the rail as they staggered forward. (Reprinted with
permission from Ref 64. Copyright 1997 American Psychological
Association (APA) and Reprinted with permission from Ref 65.
Copyright 2005 Wiley)

FI GU RE 3 | Variations in infants’ crawling patterns. Left column
shows four different crawling styles: ‘army’ crawling with the abdomen
continually resting on the ground, ‘inchworm’ crawling with the belly
on and off the ground during each cycle, standard hands-and-knees
crawling, and hands-and-feet ‘bear’ crawling. Center column
represents combinations of arms and legs used to propel the body.
Right column shows combinations of belly, hands, knees, and feet
used to maintain balance. Each row shows a unique crawling pattern.
(Reprinted with permission from Ref 58. Copyright 1998 Wiley)
consequence, most infants quickly settle into a rela-
tively stable, near-trot gait pattern.58,61 But they also
crawl on hands and feet, and combine hands, knees,
feet, and buttocks into various forms of hitching and
bum-shuffling positions that blur the line between sit-
ting and crawling.40 Balance constraints are more
severe while upright, but learning to walk is likewise
an exercise in creative problem solving with various
falling, twisting, and stepping strategies for inducing
enough disequilibrium to take steps but not so much
loss of stability to cause a fall.43,62,63
Generating new forms of locomotion can
involve cognitive skills such as means-ends problem
solving, representing goals and spatial locations, and
tool use. As illustrated in Figure 4A, when con-
fronted with challenging obstacles such as steep
slopes, cliffs, and stairs, infants search for alternative
means of descent—scooting, crawling, sliding, and
backing strategies.64,66,67 Backing is most difficult
because it requires infants to initially turn away from
the goal, coordinate backward movements, and steer
without visual guidance. On narrow bridges, infants
use a sturdy wooden handrail as a tool to augment
their balance, but they reject the handrail if it is too
far from the bridge.65,68,69 With only a wobbly rub-
ber handrail for support, they test the potential utility
of the rail, and invent various strategies for distribut-
ing body weight over the bridge and handrail
(Figure 4B).
Learning to Walk
Infants take their first walking steps at 12 months,
on average,19 but like all motor milestones, onset
ages have a wide range (8–18 months). Walking
onset awaits sufficient strength and balance to sup-
port the body on one leg as the other leg swings for-
ward.43,70,71 Both experimental and cross-cultural
studies show that experience standing, stepping, and
moving upright facilitates gains in strength and bal-
ance and accelerates the onset of walking.4,7,40 A few
minutes of daily practice with upright stepping causes
infants to begin walking weeks earlier than infants
who receive only passive exercise.55,56 Similarly, in
Caribbean and African cultures where parents delib-
erately exercise their infants’ upright skills as part of
daily massage and bathing routines (Figure 5),

© 2016 Wiley Periodicals, Inc.

 Primer wires.wiley.com/cogsci

(a) (b) (c)

(d) (e) (f)

F I G U R E 5 | Formal massage and exercise routines used in Africa, India, and the Caribbean that facilitate motor development. (a) Massage;
(b and c) Suspending the infant from the arms and feet; (d) Mother providing sitting practice; (e and f ) Practicing stepping in an upright posture.
(Reprinted with permission from Ref 72. Copyright 1988 APA and Reprinted with permission from Ref 73. Copyright 2015 Taylor and Francis)

infants walk sooner than those from the same ethnic
backgrounds who do not receive practice.16
Infants’ first steps are wobbly and uneven, with
a wide side-to-side distance between feet, a small
front-to-back distance between steps, long periods
when both feet are on the floor, and short periods
when one foot is in the air.74,75 But soon the base of
support narrows, step length increases, double sup-
port periods decrease, and infants are racing across
the floor. The steep developmental trajectory for
walking resembles the negatively accelerated perfor-
mance curves characteristic of most motor learning
tasks. Initial rapid improvements in the first 3–6
months of walking reflect infants’ discovery of the
relevant parameters that control upright balance and
propulsion.74,76–78 A protracted tapering-off period
ending between 5–7 years of age reflects subtle fine-
tuning of gait parameters.70,79 Practice, not merely
maturation, underlies improvements,74 and infants
accumulate immense amount of practice. In 1 hour
of free play, the average toddler takes about 2400
steps, travels the length of about 8 U.S. football
fields, and falls 17 times.80
Sufficient practice can lead to improvements in
endurance, strength, coordination, and balance far
beyond the norm for Western walkers.4,40 Tarahu-
maran children engage in long-distance running as
part of daily activity. As a consequence, endurance
running exceeds the abilities of most Western ultra-
marathoners: Tarahumaran children routinely run
10–40 km in a few hours and adults race
150–300 km over 24–48 hours.81 From childhood,
East African women and Nepalese porters of both
genders carry prodigious loads on their heads. Adults
in these cultures have learned to alter the biomechan-
ics of gait so as to carry loads greater than their body
weight with reduced energetic cost.82,83
Obstacle Navigation
Perception-action coupling makes locomotion func-
tional. To navigate the everyday cluttered environ-
ment, children must select the appropriate
movements and modify them accordingly, whether
crawling, walking, or riding a bicycle.84 Children
generate the requisite perceptual information through
exploratory movements—looking, touching, and test-
ing various options.7,64,85 The first studies of obstacle
navigation tested infants on a ‘visual cliff,’ a drop-off
covered in safety glass.86 But infants in such studies
can feel the glass and, after one trial, they learn that
the drop-off is only illusory—and so they cross.87
As shown in Figure 6, recent researchers have used
real cliffs, bridges, waterbeds, foam pits, water
pits, slippery surfaces, barriers, apertures, monkey
bars, car-filled streets, and other obstacles to test
the development of prospective control of
locomotion.66,67,84,88–94 Because visual and haptic
information are not in conflict on these appara-
tuses, children can be tested in dozens of trials
(an experimenter follows alongside to ensure their
safety). Many of the apparatuses are adjustable,

© 2016 Wiley Periodicals, Inc.

 WIREs Cognitive Science
FI GU RE 6 | (a) ‘Visual cliff’ with safety glass covering an
apparent drop-off. (c) Real cliff with adjustable height of drop-off.
(c) Sitting at the edge of an adjustable gap. (d) Cruising an adjustable
gap in the handrail. (e) Walking across adjustable bridges. (f )
Crawling down an adjustable slope. (g) Walking down a slope with a
Teflon-coated section. (Reprinted with permission from Ref 73.
Copyright 2015 Taylor and Francis)
allowing precise assessment of children’s ability to
gauge possibilities for locomotion.
Prelocomotor infants are sensitive to visual flow
for heading95 and depth information for a drop-
off,96 but sensitivity is not enough. Mobile infants
must learn to navigate. In their first weeks after
acquiring a new posture—sitting, crawling, cruising,
and walking—infants plunge repeatedly over the
edge of impossibly steep slopes, high cliffs, and wide
gaps. Over weeks of experience with each posture,
judgments improve so that infants attempt safe incre-
ments within their ability and avoid risky obstacles
beyond their ability.7,40
A surprising finding is that learning does not
transfer from earlier to later developing postures.
The same infants who perceive that a large gap
© 2016 Wiley Periodicals, Inc.
The development of motor behavior
precludes scooting or leaning over the edge in an
experienced sitting posture will repeatedly attempt to
crawl over the gap in a novice crawling posture.97
The same infants who perceive that a large cliff or
slope is impossible to descend in an experienced
crawling posture will repeatedly attempt to walk
when tested in a novice walking posture.64,66,67 In a
cruising posture, prewalking infants perceive pre-
cisely how large of a gap they can span with their
arms, but not with their legs.14 Clearly, failure to
transfer from earlier to later developing postures is
not due to fear of heights because the gaps, slopes,
and drop-offs are high above the ground in every
posture.98 Moreover, infants are not learning fixed
facts about the environment or their abilities because
possibilities for action change from week to week as
locomotor skills improve. Instead, infants are learn-
ing to generate and use perceptual information about
the current status of their body relative to the envi-
ronment.7,40 They are learning the relevant para-
meters for each new posture in development and the
relevant exploratory behaviors for calibrating those
parameters in a new situation.
Summary: Locomotion
Fetuses and neonates can produce leg and arm move-
ments that grossly resemble locomotion, but locomo-
tion is not hardwired or reflexive. Instead locomotor
development is tremendously plastic and responsive
to caregiving practices. And locomotion is wildly cre-
ative. Every infant discovers a unique solution for
their first crawling, walking, bum shuffling, or rolling
‘steps.’ And then they must learn to generate infor-
mation for perception and cognition to find the right
solution to suit the local constraints of the cluttered,
obstacle strewn everyday environment.
MANUAL ACTION
Manual actions begin prenatally, but outside the
womb, infants require a stable postural base to sup-
port arm movements and perceptual information to
guide movements adaptively. Tools extend children’s
manual abilities.7,99,100
Spontaneous Motility
Like all actions, manual actions appear long before
birth. Ten-week-old fetuses flex and extend their
arms, wiggle their fingers, and clench their
fists.101,102 By 14 weeks, fetuses manually explore
their own bodies, the umbilical cord, and the surface
of the uterine wall.103 By 16 weeks, fetuses bring
 Primer
hand to mouth to suck their thumbs.104 Even these
early actions are perceptually guided and planned:
Infants open their mouths in anticipation of, not in
reaction to, the arrival of their thumb.105
Spontaneous arm and hand movements con-
tinue after birth. Throughout the first year, infants
flap their arms, rotate their hands, and wiggle their
fingers, and exhibit bouts of rhythmical waving, rub-
bing, and banging while holding objects.51,106,107
Ironically, such so-called stereotypies may not be
stereotyped at all. Infants’ first banging movements
are highly variable in terms of arm trajectory. Bang-
ing becomes increasingly uniform with the arm
repeatedly tracing the same upward and downward
path.100,108
Reaching and Grasping
As in locomotion, the contextual influences of
infants’ bodies, physical environment, and social/cul-
tural environment affect the development of manual
skills. And individual infants forge their own devel-
opmental trajectories. Some infants learn to reach in
the context of spontaneous arm flaps; they stiffen the
arm joints to dampen inertial forces and direct the
arm more in the direction of the target.109 Other
infants learn to reach more conventionally, by
powering up their stationary arms in the presence of
a target. In both cases, initial attempts are usually
unsuccessful.
Goal-directed reaching requires perceptual
information about the location of the object vis-a-vis
the hand. Given appropriate postural support, neo-
nates and young infants show precursors of visually
guided reaching, extending their arms more fre-
quently while looking at a toy.110,111 Successful toy
contacts appear between 11 and 24 weeks of
age,112–114 but reaches are jerky and crooked; the
arm speeds up, slows down, and changes direction
several times prior to contact. It takes years before
children’s reaches become as smooth and straight as
those of adults.112,115 Infants ‘reach’ with their feet
at a slightly younger age than they reach with their
hands (Figure 7), showing that cephalocaudal (head
to feet) development is only a rule of thumb, not an
obligatory law of development.116
At about the same age that infants contact sta-
tionary targets, they show evidence of prospective
control in dynamic situations by intercepting moving
targets. As the toy moves along a horizontal path,
infants time their arm movements so that their hand
arrives at the location where the object will be, rather
than where it was at the start of the reach.117–119 Vis-
ual information for frontally approaching targets is
© 2016 Wiley Periodicals, Inc.
wires.wiley.com/cogsci
F I G U R E 7 | Three-month-old infant ‘feet reaching’ by contacting
an object with the foot. (Reprinted with permission from Ref 73.
Copyright 2015 Taylor and Francis)
different: The toy expands in the field of view. By
8–9 months of age, infants precisely gauge whether
balls approaching at different speeds are catchable
and they initiate interceptive arm movements based
on visual information for time to contact.120
Older children and adults rely on view of the
hand as well as view of the target to guide reach-
ing.121,122 However, young infants do not benefit
from being able to see their moving hand. Research-
ers can measure the importance of continual visual
feedback by turning off the lights once a reach begins
(the toy glows to mark its location) or by occluding
sight of the hand and arm with a cloth barrier.
Infants begin reaching for objects in the light and
dark at the same age, and are equally successful in
both conditions.114 Moreover, the kinematics of
infants’ reaching trajectories in the light and dark are
indistinguishable123 and do not require sight of the
hand.124 In other words, infants’ zigzag reach trajec-
tories do not necessarily mean that they visually
track their hand because infants display equally jerky
reaches when they cannot see their hand. Jerky tra-
jectories may result in part from postural con-
straints17 and unanticipated reactive forces.112
Much to infants’ frustration, getting the hand
to the right place is only part of the problem.
Reaching precedes grasping because control of the
arms precedes control of the hands. Normally,
3-month-olds merely swat at objects because they
lack the requisite hand control for grasping and do
not use perceptual information about object proper-
ties to plan the grasp. However, with the help of
sticky Velcro mittens and Velcro covered toys, swats
 WIREs Cognitive Science
are sufficient to pick up an object, thereby allowing
‘pre-graspers’ to reap some of the benefits of grasp-
ing objects that only older infants normally experi-
ence.125 These early benefits have both immediate
and long-lasting effects on manual skill.126–128 With
increased hand/finger control, infants adapt their
grip configuration to object properties, but they do
so after contacting the object, not prospectively dur-
ing the reach.129 Prospective control of grasping
based on visual information for object size, orienta-
tion, and substance appears months after infants
begin reaching.130–133
Exploring Objects
An object in hand opens up new opportunities for
visual, manual, and oral exploration, and with
increasing skill, object exploration becomes increas-
ingly multimodal.134,135 At first, infants use their
hands only to bring objects up to the face for looking
and mouthing.136 Increased grip strength allows
infants to alternate between looking and mouthing,
providing multimodal information about object
properties. Soon, manual skills progress beyond mere
holding. Infants heft, rub, squeeze, and finger
objects.31,135,137 Later, infants coordinate visual and
manual exploration by transferring objects from
hand to hand and rotating them in front of their
(a)
(d)
Both
hands
FI GU RE 8 | Nine-month-old infant grasping a spoon (a) by the bowl or (b) with an ulnar grip that points the bowl away from the mouth.
(c) An 18-month-old using a radial grip that correctly brings food to the mouth. (d) Variety of pen grips used by 3- and 5-year-olds and adults.
(Reprinted with permission from Ref 143. Copyright 1999 APA and Reprinted with permission from Ref 144. Copyright 1998 Wiley)
The development of motor behavior
eyes.25 Hands begin to serve complementary func-
tions, one supporting the object and keeping it in
view, the other generating information about object
properties by fingering or palpating.138 Infants
explore the relations between object and surface
properties by banging a hard block against a rigid
surface to make a noise or rubbing a soft block
against the surface.107
Extending Abilities with Tools
Tool use has its roots in early motor actions and
relies on motor actions for its execution.99,100,108
Young infants’ spontaneous banging and rubbing
become preschoolers’ hammering and drawing. Fetal
hand-to-mouth behaviors become self-feeding with a
spoon. Exploring relations between objects and sur-
faces sets the stage for using objects as effective tools.
Tool use requires infants to perceive that a goal
is beyond their abilities, recognize that an object can
serve as a means to augment their abilities, and exe-
cute the necessary movements to use the tool. Each
of these steps in real time must first be acquired in
development. For example, very young infants per-
ceive when an object is out of reach.139 Months later,
they use hooks, canes, and rakes to acquire out-of-
reach objects, but only if the target object is already
placed inside the crook of the tool.140–142 And still
(b) (c)
© 2016 Wiley Periodicals, Inc.
 Primer
later they perceive the full implications of the spatial
relations by orienting the tools to place the target in
the crook. Observing caregivers or other adults use a
tool effectively provides a powerful impetus for
learning142
Implementation often stands in the way of func-
tional tool use. Nine-month-olds grasp a spoon filled
with applesauce by the bowl end rather than by the
handle (getting a handful of applesauce), or with a
grip that points the food away from the mouth so
that they cannot eat (Figure 8A,B). Eighteen-month-
olds perceive the optimal grasp for delivering food to
their own mouth and plan their grasp prospectively,
but their planning is less efficient when feeding a doll
(Figure 8C).143,145,146 Two-year-olds adapt their
grasp to use a spoon with a bent handle.147 But even
4-year-olds fail to realize that they must use an
underhand grip to grasp a spoon or hammer pointing
away from their dominant hand.148 Implementing a
writing or drawing instrument poses similar pro-
blems for older children.144 Three-year-olds use
11 different grip configurations to draw straight lines
(including using both hands to hold the pen) and
individual children vary their grips from trial to trial
(Figure 8D). Variability decreases by 5 years of age
when most children begin formal schooling, and chil-
dren converge on one of the two common adult
grips.
Summary: Manual Action
Beginning prenatally, manual actions are perceptu-
ally guided and serve exploratory functions. Many of
infants’ spontaneous arm and hand movements are
co-opted for goal-directed manual actions and tool
use. Infants use vision to locate the target of a reach
and to preshape their hand for grasping, but they do
not require sight of their hand to get it to a target.
Exploring objects is a multimodal activity involving
eyes, hands, fingers, and mouth. Boosting up manual
skills can jump start the cascade of opportunities for
learning.
FACIAL ACTION
All the parts of the face begin moving prenatally,
including the eyes while they are still fused shut.
After birth, infants continue to produce spontaneous
facial movements, but facial actions become integral
to everyday function. The simple ability to swallow
is critical for suckling, eating, and talking. Vocaliza-
tions and facial expressions are fundamental for com-
munication. Head and eye movements provide the
basis for visual exploration of the environment.
© 2016 Wiley Periodicals, Inc.
wires.wiley.com/cogsci
Swallowing, Sucking, and Chewing
Actions like swallowing are normally so innocuous
that we do not recognize the tremendous coordina-
tion required. Fetuses make swallowing, sucking, and
breathing movements, but since they do not breathe
air or eat, the movements are not coordinated.149
However, to nurse without choking or swallowing
air, newborns must coordinate movements of tongue,
jaws, and lips to create suction, draw liquid into the
mouth, pull the liquid into the pharynx, and divert
the liquid to the esophagus while pulling air into the
trachea.150–152 Infants solve the timing problem by
coordinating suck–swallow–respiration patterns at a
ratio of 1:1:1 or 2:2:1.153–155
Chewing solid food is more complicated
because the food must be masticated before it can be
swallowed. Newborns can mush up a small piece of
banana and move it around the mouth with jaws and
tongue.156 However, infants rely on lateral jaw
movements to do most of the work of chewing,
whereas older children and adults use rotary jaw
movements and incorporate more prospective actions
of the lips and tongue.157 Infants produce the same
chewing movements regardless of the type of food,
whereas older children select appropriate jaw move-
ments and muscle forces based on the food
consistency.157
Facial Gestures and Speech
Facial expressions and vocalizations appear long
before infants can convey feelings and communicate
ideas. Fetuses produce smiles, grimaces, and facial
movements that resemble adult-like expressions of
laughter, crying, and pain.158,159 Neonates produce
characteristic facial gestures to strong stimuli such as
nose wrinkling and furrowed brows to noxious
smells.160 Newborns smile most while asleep, about
one smile every 5 min.161,162 Awake infants begin to
display social smiles and laughter by 2–5 months of
age while gazing at caregivers or in response to posi-
tive stimulation.163 Perhaps because they are so criti-
cal for social interaction, facial expressions are
highly redundant so that muscles distributed
throughout the face work in concert; eyebrows can
convey basic facial expressions as effectively as the
mouth. In fact, infants who lack the ability to move
specific parts of their faces due to severe craniofacial
anomalies, cleft lip/palate, or hemangiomas produce
recognizable smiles, cries, and interested
expressions.164
The movements needed for speech production
are perhaps the most complex movements children
learn.165 The jaws, lips, and tongue must be
 WIREs Cognitive Science
precisely positioned to shape each sound as air tra-
vels through the oral and nasal cavities. Both speed
and accuracy are major challenges in speech devel-
opment. Adult-like speech is incredibly fast, encom-
passing up to 15 sounds per second.165 As in the
development of chewing, infants discover functional
strategies to produce speech sounds, but their move-
ments are not adult-like. For example, adults use
quick simultaneous movements of the jaws and lips
to babble (baba, mama), whereas infants rely prima-
rily on jaw movements, which are easier to con-
trol.166 Between 2–6 years of age, children gain
better control over their lips and incorporate those
movements into the previously established jaw
movements, allowing them to produce a greater
variety of speech sounds.150
Looking
Visual perception involves pointing the eyes in the
right direction. But looking usually involves more
than moving the eyes. It involves coordination
among body, head, and eyes to bring a desired loca-
tion into view.167 For newborns who can barely
turn their heads, the control problem is simplified:
They typically watch whatever happens to be in
front of them, whether faces, hands, objects, or
more complex scenes.168–170 Even after posture
improves and infants can sit, crawl, and walk, much
of what they see is opportunistic. Toddlers are less
likely to tilt their head up to look at mother’s face
than to point their gaze straight ahead at her knees;
they see what’s in their hands or someone else’s
hands because the hands are already in their field of
view.23,171–173
Like other motor actions, looking is more func-
tional and adaptive when eye, head, and body move-
ments are controlled prospectively. To track a
moving object, infants must anticipate its speed and
trajectory to keep their eyes moving at the right pace.
Large targets moving in predictable ways are easier
for young infants to smoothly pursue with their
eyes.174 When the target moves too quickly, the eyes
lag behind, so infants often make corrective saccades
to catch up to the target. Over months of practice,
infants track smaller targets at faster speeds,175
resorting less often to corrective saccades. Initially,
infants keep their eyes on the target, but their head
lags behind. By 4–5 months, infants coordinate
movements of eyes and head to smoothly follow
moving objects.176 While watching an object move
repeatedly behind an occluder, 4-month-olds keep
their eyes on the point where the object disappeared
and then struggle to catch up to its motion after it
© 2016 Wiley Periodicals, Inc.
The development of motor behavior
(a)
Scene
camera
Eye
Wireless
camera transmitter
(b)
FI G URE 9 | (a) Head-mounted eye-tracker worn by a 14-month-old
infant. An outward facing ‘scene camera’ records the infant’s field of
view, and an inward facing ‘eye camera’ records movements of the
infant’s right eye. Computer software calculates point of gaze.
(b) Processed gaze video with red crosshair showing the infant’s point of
gaze. (Reprinted with permission from Ref 171. Copyright 2011 Wiley)
appears on the other side; 6-month-olds visually
anticipate the location where the object will reap-
pear, indicating their understanding that the object
exists even when it is out of sight.177
As in the case of walking, infants amass tre-
mendous amounts of experience while learning to
look. In 1 day, infants shift their gaze roughly
50,000 times.178 By 2 months of age, infants have
accumulated 200 hours of visual experience177 and
by 3.5 months of age, researchers estimate that
infants have produced 3–6 million eye movements.179
However, researchers know very little about what
infants actually see outside of the laboratory. Head-
mounted eye tracking provides a new method for
observing infants’ eye movements during uncon-
strained, spontaneous activity. As shown in Figure 9,
infants wear two small cameras that record their eye
movements and field of view for real-time gaze pro-
cessing. Toddlers seamlessly distribute visual atten-
tion among multiple motor tasks, looking toward
 Primer wires.wiley.com/cogsci

obstacles to guide crawling and walking, fixating CONCLUSION

objects to guide the hand while reaching, and glanc-
ing occasionally at caregivers to initiate or respond
to social interactions.171 Visual guidance becomes
increasingly efficient over the course of development.
Quick glances toward obstacles from a distance elicit
more costly types of exploration such as touching.85
Infants’ short bodies serendipitously contribute to
successful obstacle navigation because their field of
view includes more of the floor compared with older
children and adults, who primarily guide locomotion
with visual information from the periphery.171,180
Summary: Facial Action
Facial actions include many of our most prized and
basic social skills—talking, facial gestures, eating and
drinking, and looking at others and at the environ-
ment. And each of these skills sets off a new cascade
of interactions. Infants’ solutions for moving the vari-
ous parts of their face often differ from those of
adults, but they get the job done in that developmen-
tal niche.
The study of motor development is really the study of
behavioral development. As such, it can provide a use-
ful window into general processes of development
because the topic of study—movement—is directly
observable. Researchers in motor development have
always recognized the importance of the bodily con-
text.7 How could they do otherwise? Movements
depend on physical forces and the moment-to-moment
changes and developmental status of the body affect
forces. The developmental systems perspective
encourages researchers also to consider a larger con-
text that includes the physical and social/cultural envi-
ronment, and to view motor behaviors as potentially
both cause and consequence of developmental change
in other psychological domains (see Goldin-Meadow,
Using our hands to change our minds, WIREs Cogn
Sci, also in the collection How We Develop).
Although prominent developmental theorists have
long recognized the importance of motor development
for psychological development more generally,5,6 only
recently have researchers begun to systematically map
out these developmental pathways.

ACKNOWLEDGMENTS
The writing of this article was supported by NICHD Grant R37-HD33486 to K. E. Adolph.

FURTHER READING
Adolph KE, Berger SE. Motor development. In: Kuhn D, Siegler RS, eds. Handbook of Child Psychology: Cognition, Per-
ception, and Language, vol. 2. 6th ed. New York: Wiley; 2006, 161–213.
Gibson EJ, Pick AD. An Ecological Approach to Perceptual Learning and Development. New York: Oxford University
Press; 2000.
Smitsman AW, Corbetta D. Action in infancy: perspectives, concepts, and challenges. In: Bremner JG, Wachs TD, eds. The
Wiley-Blackwell Handbook of Infant Development. 2nd ed. Chichester and West Sussex: Wiley-Blackwell Ltd; 2010,
167–203.
Thelen E. Motor development: a new synthesis. Am Psychol 1995, 50:79–95.

REFERENCES
1. Bernstein NA. On dexterity and its development. In: 4. Adolph KE, Karasik LB, Tamis-LeMonda CS. Motor
Latash ML, Turvey MT, eds. Dexterity and Its Devel- skills. In: Bornstein MH, ed. Handbook of Cultural
opment. Mahwah, NJ: Erlbaum; 1996, 3–244. Development Science. Domains of Development across
Cultures, vol. 1. New York: Taylor and Francis;
2. Gibson EJ. Has psychology a future? Psychol Sci 1994,
2010, 61–88.
5:69–76.
5. Gibson EJ. Exploratory behavior in the development
3. Keen R. The development of problem solving in young
of perceiving, acting, and the acquiring of knowledge.
children: a critical cognitive skill. Annu Rev Psychol
Annu Rev Psychol 1988, 39:1–41.
2011, 62:1–21.

© 2016 Wiley Periodicals, Inc.

 WIREs Cognitive Science
6. Piaget J. The Construction of Reality in the Child.
New York: Basic Books; 1954.
7. Adolph KE, Robinson SR. Motor development. In:
Liben L, Muller U, eds. Handbook of Child Psy-
chology and Developmental Science. Cognitive Pro-
cesses, vol. 2. 7th ed. New York: Wiley; 2015,
114–157.
8. Thelen E. Dynamic systems theory and the complex-
ity of change. Psychoanal Dialogues 2005,
15:255–283.
9. Campos JJ, Anderson DI, Barbu-Roth MA,
Hubbard EM, Hertenstein MJ, Witherington DC.
Travel broadens the mind. Infancy 2000, 1:149–219.
10. Reed ES. An outline of a theory of action systems.
J Mot Behav 1982, 14:98–134.
11. D’Elia A, Pighetti M, Moccia G, Santangelo N. Spon-
taneous motor activity in normal fetuses. Early Hum
Dev 2001, 65:139–147.
12. de Vries JIP, Hopkins B. Fetal movements and pos-
tures: what do they mean for postnatal development?
In: Hopkins B, Johnson SP, eds. Prenatal Development
of Postnatal Functions. Westport, CT: Praeger; 2005,
177–220.
13. Gesell A. The ontogenesis of infant behavior. In:
Carmichael L, ed. Manual of Child Psychology.
New York: John Wiley; 1946, 295–331.
14. Adolph KE, Berger SE, Leo AJ. Developmental conti-
nuity? Crawling, cruising, and walking. Dev Sci 2011,
14:306–318.
15. Atun-Einy O, Berger SE, Scher A. Pulling to stand:
common trajectories and individual differences. Dev
Psychobiol 2012, 54:187–198.
16. Hopkins B, Westra T. Motor development, maternal
expectations, and the role of handling. Infant Behav
Dev 1990, 13:117–122.
17. Hopkins B, Ronnqvist L. Facilitating postural control:
effects on the reaching behavior of 6-month-old
infants. Dev Psychobiol 2002, 40:168–182.
18. Saavedra SL, van Donkelaar P, Woollacott MH.
Learning about gravity: segmental assessment of
upright control as infants develop independent sitting.
J Neurophysiol 2012, 108:2215–2229.
19. Martorell R, Onis M, Martines J, Black M,
Onyango A, Dewey KG. WHO motor development
study: windows of achievement for six gross motor
development milestones. Acta Paediatr 2006,
95(S450):86–95.
20. Karasik LB, Tamis-LeMonda CS, Adolph KE,
Bornstein MH. Places and postures: a cross-cultural
comparison of sitting in 5-month-olds. J Cross Cult
Psychol 2015, 46:1023–1038.
21. Kretch KS, Adolph KE. Active vision in passive loco-
motion: real-world free viewing in infants and adults.
Dev Sci 2015, 18:736–750.
© 2016 Wiley Periodicals, Inc.
The development of motor behavior
22. Fogel A, Messinger DS, Dickson KL, Hsu H. Posture
and gaze in early mother-infant communication: syn-
chronization of developmental trajectories. Dev Sci
1999, 2:325–332.
23. Kretch KS, Franchak JM, Adolph KE. Crawling and
walking infants see the world differently. Child Devel-
opment 2014, 85:1503–1518.
24. Rachwani J, Santamaria V, Saavedra SL, Wood S,
Porter F, Woollacott MH. Segmental trunk control
acquisition and reaching in typically developing
infants. Exp Brain Res 2013, 228:131–139.
25. Soska KC, Adolph KE. Postural position constrains
multimodal object exploration in infants. Infancy
2014, 19:138–161.
26. Harbourne RT, Lobo MA, Karst GM, Galloway JC.
Sit happens: does sitting development perturb reaching
development, or vice versa? Infant Behav Dev 2013,
36:438–450.
27. Carvalho RP, Tudella E, Calijouw SR,
Savelsbergh GJP. Early control of reaching: effects of
experience and body orientation. Infant Behav Dev
2008, 31:23–33.
28. Soska KC, Adolph KE, Johnson SP. Systems in devel-
opment: motor skill acquisition facilitates three-
dimensional object completion. Dev Psychol 2010,
46:129–138.
29. Baumgartner HA, Oakes LM. Investigating the rela-
tion between infants’ manual activity with objects and
their perception of dynamic events. Infancy 2013,
18:983–1006.
30. Libertus K, Gibson J, Hidayatallah NZ, Hirtle J,
Adcock RA, Needham AW. Size matters: how age and
reaching experience shape infants’ preferences for dif-
ferent sized objects. Infant Behav Dev 2013,
36:189–198.
31. Eppler MA. Development of manipulatory skills and
the deployment of attention. Infant Behav Dev 1995,
18:391–405.
32. Loucks J, Sommerville JA. The role of motor experi-
ence in understanding action function: the case of the
precision grasp. Child Dev 2012, 83:801–809.
33. Daum MM, Prinz W, Aschersleben G. Perception and
production of object-related grasping in 6-month-olds.
J Exp Child Psychol 2011, 108:810–818.
34. Deffeyes JE, Harbourne RT, Stuberg WA, Stergiou N.
Approximate entropy used to assess sitting postural
sway of infants with developmental delay. Infant
Behav Dev 2011, 34:81–99.
35. Metcalfe JS, Chen LC, Chang TY, McDowell K,
Jeka JJ, Clark JE. The temporal organization of pos-
ture changes during the first year of independent walk-
ing. Exp Brain Res 2005, 161:405–416.
36. Chen LC, Metcalfe JS, Chang TY, Jeka JJ, Clark JE.
The development of infant upright posture: sway less
 Primer
or sway differently? Exp Brain Res 2008,
186:293–303.
37. Bertenthal BI, Bai DL. Infants’ sensitivity to optical
flow for controlling posture. Dev Psychol 1989,
25:936–945.
38. Dahl A, Campos JJ, Anderson DI, Uchiyama I,
Witherington DC, Ueno M, Poutrain-Lejeune L,
Barbu-Roth M. The epigenesis of wariness of heights.
Psychol Sci 2013, 24:1361–1367.
39. Chen LC, Jeka J, Clark JE. Development of adaptive
sensorimotor control in infant sitting posture. Gait
Posture 2016, 45:157–163.
40. Adolph KE, Robinson SR. The road to walking: what
learning to walk tells us about development. In:
Zelazo P, ed. Oxford Handbook of Developmental
Psychology. New York: Oxford University Press;
2013, 403–443.
41. Andre-Thomas M, Autgaerden S. Locomotion from
Pre- to Post-Natal Life. Lavenham, Suffolk: Spastics
Society Medical Education and Information Unit in
association with William Heinemann Medical Books,
Ltd.; 1966.
42. Forssberg H. Ontogeny of human locomotor control.
I. Infant stepping, supported locomotion, and transi-
tion to independent locomotion. Exp Brain Res 1985,
57:480–493.
43. McGraw MB. The Neuromuscular Maturation of the
Human Infant. New York: Columbia University
Press; 1945.
44. Dominici N, Ivanenko YP, Cappellini G, d’Avella A,
Mondi V, Cicchese M, Fabiano A, Silei T, Di Paolo A,
Giannini C, et al. Locomotor primitives in newborn
babies and their development. Science 2011,
334:997–999.
45. Barbu-Roth M, Anderson DI, Despres A, Provasi J,
Cabrol D, Campos JJ. Neonatal stepping in relation to
terrestrial optic flow. Child Dev 2009, 80:8–14.
46. Barbu-Roth M, Anderson DI, Despres A, Streeter RJ,
Caborl D, Trukillo M, Campos JJ, Provasi J. Air step-
ping in response to optic flows that move toward and
away from the neonate. Dev Psychobiol 2014,
56:1142–1149.
47. Angulo-Kinzler RM, Ulrich BD, Thelen E. Three-
month-old infants can select specific leg motor solu-
tions. Motor Control 2002, 6:52–68.
48. Rovee-Collier CK, Morrongiello BA, Aron M,
Kupersmidt J. Topographical response differentiation
and reversal in 3-month-old infants. Infant Behav Dev
1978, 1:323–333.
49. Thelen E. Three-month-old infants can learn task-
specific patterns of interlimb coordination. Psychol Sci
1994, 5:280–285.
50. Thelen E, Fisher DM. Newborn stepping: an explana-
tion for a “disappearing reflex”. Dev Psychol 1982,
18:760–775.
© 2016 Wiley Periodicals, Inc.
wires.wiley.com/cogsci
51. Thelen E. Rhythmical stereotypies in normal human
infants. Anim Behav 1979, 27:699–715.
52. Thelen E. Treadmill-elicited stepping in seven-month-
old infants. Child Dev 1986, 57:1498–1506.
53. Yang JF, Lam T, Pang MYC, Lamont E,
Musselman K, Seinen E. Infant stepping: a window to
the behavior of the human pattern generator for walk-
ing. Can J Physiol Pharmacol 2004, 82:662–674.
54. Thelen E, Fisher DM, Ridley-Johnson R. The relation-
ship between physical growth and a newborn reflex.
Infant Behav Dev 1984, 7:479–493.
55. Zelazo PR. The development of walking: new findings
on old assumptions. J Mot Behav 1983, 2:99–137.
56. Zelazo PR, Zelazo NA, Kolb S. “Walking” in the new-
born. Science 1972, 176:314–315.
57. van der Meer ALH, Ramstad M, van der Weel FR.
Choosing the shortest way to mum: auditory guided
rotation in 6- to 9-moth-old infants. Infant Behav Dev
2008, 31:207–216.
58. Adolph KE, Vereijken B, Denny MA. Learning to
crawl. Child Dev 1998, 69:1299–1312.
59. Patrick SK, Noah JA, Yang JF. Developmental con-
straints of quadrupedal coordination across crawling
styles in human infants. J Neurophysiol 2012,
107:3050–3061.
60. Majnemer A, Barr RG. Influence of supine sleep posi-
tioning on early motor milestone acquisition. Dev Med
Child Neurol 2005, 47:370–376.
61. Patrick SK, Noah JA, Yang JF. Interlimb coordination
in human crawling reveals similarities in development
and neural control with quadrupeds. J Neurophysiol
2009, 101:603–613.
62. McCollum G, Holroyd C, Castelfranco AM. Forms of
early walking. J Theor Biol 1995, 176:373–390.
63. Snapp-Childs W, Corbetta D. Evidence of early strate-
gies in learning to walk. Infancy 2009, 14:101–116.
64. Adolph KE. Learning in the development of infant
locomotion. Monogr Soc Res Child Dev 1997, 62.
65. Berger SE, Adolph KE, Lobo SA. Out of the toolbox:
toddlers differentiate wobbly and wooden handrails.
Child Dev 2005, 76:1294–1307.
66. Kretch KS, Adolph KE. Cliff or step? Posture-specific
learning at the edge of a drop-off. Child Dev 2013,
84:226–240.
67. Karasik LB, Tamis-LeMonda CS, Adolph KE. Deci-
sions at the brink: locomotor experience affects
infants’ use of social information on an adjustable
drop-off. Front Psychol 2016, 7:797.
68. Berger SE, Adolph KE. Infants use handrails as tools in
a locomotor task. Dev Psychol 2003, 39:594–605.
69. Berger SE, Adolph KE, Kavookjian AE. Bridging the
gap: solving spatial means-ends relations in a locomo-
tor task. Child Dev 2010, 81:1367–1375.
 WIREs Cognitive Science
70. Bril B, Ledebt A. Head coordination as a means to
assist sensory integration in learning to walk. Neurosci
Biobehav Rev 1998, 22:555–563.
71. Hallemans A, De Clercq D, Aerts P. Changes in 3D
joint dynamics during the first 5 months after the onset
of independent walking: a longitudinal follow-up
study. Gait Posture 2006, 24:270–279.
72. Hopkins B, Westra T. Maternal handling and motor
development: an intracultural study. Genet Soc Gen
Psychol Monogr 1988, 114:379–408.
73. Adolph KE, Berger SE. Physical and motor develop-
ment. In: Bornstein MH, Lamb ME, eds. Development
Science: An Advanced Textbook. 7th ed. New York:
Psychology Press/Taylor & Francis; 2015, 261–333.
74. Adolph KE, Vereijken B, Shrout PE. What changes in
infant walking and why. Child Dev 2003,
74:474–497.
75. Bril B, Breniere Y. Postural requirements and progres-
sion velocity in young walkers. J Mot Behav 1992,
24:105–116.
76. Bril B, Breniere Y. Posture and independent locomo-
tion in early childhood: learning to walk or learning
dynamic postural control? In: Savelsbergh GJP,
ed. The Development of Coordination in Infancy.
North-Holland: Elsevier; 1993, 337–358.
77. Ivanenko YP, Dominici N, Cappellini G, Lacquaniti F.
Kinematics in newly walking toddlers does not depend
upon postural stability. J Neurophysiol 2005,
94:754–763.
78. Bril B, Dupuy L, Dietrich G, Corbetta D. Learning to
tune the antero-posterior propulsive forces during
walking: a necessary skill for mastering upright loco-
motion in toddlers. Exp Brain Res 2015,
233:2903–2912.
79. Sutherland DH, Olshen RA, Biden EN, Wyatt MP.
The Development of Mature Walking. London: Mac
Keith Press; 1988.
80. Adolph KE, Cole WG, Komati M, Garciaguirre JS,
Badaly D, Lingeman JM, Chan G, Sotsky RB. How do
you learn to walk? Thousands of steps and dozens of
falls per day. Psychol Sci 2012, 23:1387–1394.
81. Bennett WC, Zingg RM. The Tarahumara, an Indian
tribe of northern Mexico. Chicago, IL: University of
Chicago Press; 1935.
82. Bastien GJ, Schepens B, Willems PA, Heglund NC.
Energetics of load carrying in Nepalese porters. Science
2005, 308:1755.
83. Maloiy GMO, Heglund NC, Prager LM, Cavagna GA,
Taylor CR. Energetic costs of carrying loads: have
African women discovered an economic way? Nature
1986, 319:668–669.
84. Plumert JM, Kearney JK. How do children perceive
and act on dynamic affordances in crossing traffic-
filled roads? Child Dev Perspect 2014, 8:207–212.
© 2016 Wiley Periodicals, Inc.
The development of motor behavior
85. Kretch KS, Adolph KE. The organization of explora-
tory behaviors in infant locomotor planning. Dev Sci.
In press. doi: 10.1111/desc.12421.
86. Gibson EJ, Walk RD. The “visual cliff”. Sci Am 1960,
202:64–71.
87. Campos JJ, Hiatt S, Ramsay D, Henderson C,
Svejda M. The emergence of fear on the visual cliff. In:
Lewis M, Rosenblum L, eds. The Development of
Affect. New York: Plenum; 1978, 149–182.
88. Adolph KE, Joh AS, Eppler MA. Infants’ perception of
affordances of slopes under high and low friction con-
ditions. J Exp Psychol Hum Percept Perform 2010,
36:797–811.
89. Burnay C, Cordovil R. Crawling experience predicts
avoidance of real cliffs and water cliffs: insight from a
new paradigm. Infancy 2016, 21:677–684.
90. Franchak JM, Adolph KE. What infants know and
what they do: perceiving possibilities for walking
through openings. Dev Psychol 2012, 48:1254–1261.
91. Gibson EJ, Riccio G, Schmuckler MA, Stoffregen TA,
Rosenberg D, Taormina J. Detection of the traversabil-
ity of surfaces by crawling and walking infants. J Exp
Psychol Hum Percept Perform 1987, 13:533–544.
92. Joh AS, Adolph KE. Learning from falling. Child Dev
2006, 77:89–102.
93. Schmuckler MA. Development of visually guided loco-
motion: barrier crossing by toddlers. Ecol Psychol
1996, 8:209–236.
94. van der Meer ALH. Visual guidance of passing under
a barrier. Early Dev Parenting 1997, 6:149–157.
95. Gilmore RO, Baker TJ, Grobman KH. Stability in
young infants’ discrimination of optic flow. Dev Psy-
chol 2004, 40:259–270.
96. Campos JJ, Bertenthal BI, Kermoian R. Early experi-
ence and emotional development: the emergence of
wariness of heights. Psychol Sci 1992, 3:61–64.
97. Adolph KE. Specificity of learning: why infants fall
over a veritable cliff. Psychol Sci 2000, 11:290–295.
98. Adolph KE, Kretch KS, LoBue V. Fear of heights in
infants? Curr Dir Psychol Sci 2014, 23:60–66.
99. Smitsman AW, Bongers RM. Tool use and tool mak-
ing: a developmental action perspective. In: Valsiner J,
Connolly KJ, eds. Handbook of Developmental Psy-
chology. London: Sage; 2003, 172–193.
100. Lockman JJ, Kahrs BA. Tool using. Child Dev Per-
spect 2014, 8:231–236.
101. Prechtl HFR. Ultrasound studies of human fetal beha-
viour. Early Hum Dev 1985, 12:91–98.
102. Prechtl HFR. Prenatal motor development. In:
Wade MG, Whiting HTA, eds. Motor Development
in Children: Aspects of Coordination and Control.
Dordrecht: Martinus Nijhoff Publishing;
1986, 53–64.
 Primer
103. Sparling JW, van Tol J, Chescheir NC. Fetal and neo-
natal hand movement. Phys Ther 1999, 79:24–39.
104. Hepper PG, Shahidullah S, White R. Handedness in
the human fetus. Neuropsychologia 1991,
29:1107–1111.
105. Reissland N, Francis B, Aydin E, Mason J, Schaal B.
The development of anticipation in the fetus: a longi-
tudinal account of human fetal mouth movements in
reaction to and anticipation of touch. Dev Psychobiol
2014, 56:955–963.
106. Piek JP, Carman R. Developmental profiles of sponta-
neous movements in infants. Early Hum Dev 1994,
39:109–126.
107. Fontenelle SA, Kahrs BA, Neal SA, Newton AT,
Lockman JJ. Infant manual exploration of composite
substrates. J Exp Child Psychol 2007, 98:153–167.
108. Kahrs BA, Jung WP, Lockman JJ. Motor origins of
tool use. Child Dev 2013, 84:810–816.
109. Thelen E, Corbetta D, Kamm K, Spencer JP,
Schneider K, Zernicke RF. The transition to reaching:
mapping intention and intrinsic dynamics. Child Dev
1993, 64:1058–1098.
110. Lee HM, Bhat A, Scholz JP, Galloway JC. Toy-
oriented changes during early arm movements IV:
shoulder-elbow coordination. Infant Behav Dev
2008, 31:447–469.
111. Lee MH, Ranganathan R, Newell KM. Changes in
object-oriented arm movements that precede the tran-
sition to goal-directed reaching in infancy. Dev Psy-
chobiol 2011, 53:685–693.
112. Berthier NE, Keen RE. Development of reaching in
infancy. Exp Brain Res 2006, 169:507–518.
113. Konczak J, Dichgans J. The development toward ster-
eotypic arm kinematics during reaching in the first
3 years of life. Exp Brain Res 1997, 117:346–354.
114. Clifton RK, Muir DW, Ashmead DH, Clarkson MG.
Is visually guided reaching in early infancy a myth?
Child Dev 1993, 64:1099–1110.
115. Schneiberg S, Sveistrup H, McFadyen B, McKinley P,
Levin MF. The development of coordination of
reach-to-grasp movements in children. Exp Brain Res
2002, 146:142–154.
116. Galloway JC, Thelen E. Feet first: object exploration
in young infants. Infant Behav Dev 2004,
27:107–112.
117. von Hofsten C. Catching skills in infancy. J Exp Psy-
chol Hum Percept Perform 1983, 9:75–85.
118. von Hofsten C, Lindhagen K. Observations on the
development of reaching for moving objects. J Exp
Child Psychol 1979, 28:158–173.
119. Fagard J, Spelke ES, von Hofsten C. Reaching and
grasping a moving object in 6-, 8-, and 10-month-old
infants: laterality and performance. Infant Behav Dev
2009, 32:137–146.
© 2016 Wiley Periodicals, Inc.
wires.wiley.com/cogsci
120. van Hof P, van der Kamp J, Savelsbergh GJP. The
relation between infants’ perception of catchableness
and the control of catching. Dev Psychol 2008,
44:182–194.
121. Churchill A, Hopkins B, Ronnqvist L, Vogt S. Vision
of the hand and environmental context in human pre-
hension. Exp Brain Res 2000, 134:81–89.
122. Kuhtz-Buschbeck JP, Stolze H, Boczek-Funcke A,
Johnk K, Heinrichs H, Illert M. Kinematic analysis of
prehension movements in children. Behav Brain Res
1998, 93:131–141.
123. Clifton RK, Rochat P, Robin DJ, Berthier NE. Multi-
modal perception in the control of infant reaching.
J Exp Psychol Hum Percept Perform 1994,
20:876–886.
124. Lee MH, Newell KM. Visual feedback of hand trajec-
tory and the development of infant prehension. Infant
Behav Dev 2012, 35:273–279.
125. Needham AW, Barrett T, Peterman K. A pick-me-up
for infants’ exploratory skills: early simulated experi-
ences reaching for objects using “sticky” mittens
enhances young infants’ object exploration skills.
Infant Behav Dev 2002, 25:279–295.
126. Libertus K, Needham AW. Teach to reach: the effects
of active vs. passive reaching experiences on action
and perception. Vision Res 2010, 50:2750–2757.
127. Wiesen SE, Watkins RM, Needham AW. Active
motor training has long-term effects on infants’ object
exploration. Front Psychol 2016, 7:599.
128. Libertus K, Joh AS, Needham AW. Motor training at
3 months affects object exploration 12 months later.
Dev Sci 2015, 6:1058–1066.
129. Vollmer B, Forssberg H. Development of grasping
and object manipulation. In: Nowak DA,
Hermsdorfer J, eds. Sensorimotor Control of Grasp-
ing: Physiology and Pathophysiology. Cambridge:
Cambridge University Press; 2009, 235–249.
130. Barrett TM, Traupman E, Needham AW. Infants’ vis-
ual anticipation of object structure in grasp planning.
Infant Behav Dev 2008, 31:1–9.
131. Fagard J. Linked proximal and distal changes in the
reaching behavior of 5- to 12-month-old human
infants grasping objects of different sizes. Infant
Behav Dev 2000, 23:317–329.
132. Witherington DC. The development of prospective
grasping control between 5 and 7 months: a longitu-
dinal study. Infancy 2005, 7:143–161.
133. von Hofsten C, Johansson K. Planning to reach for a
rotating rod: developmental aspects. ZEEP 2009,
41:207–213.
134. Needham AW. Improvements in object exploration
skills may facilitate the development of object segre-
gation in early infancy. J Cognit Dev 2000,
1:131–156.
 WIREs Cognitive Science
135. Rochat P. Object manipulation and exploration in 2-
to 5-month-old infants. Dev Psychol 1989,
25:871–884.
136. Norris B, Smith S. Research into the mouthing beha-
viour of children up to 5 years old, 2002.
137. Palmer CF. The discriminating nature of infants’
exploratory actions. Dev Psychol 1989, 25:885–893.
138. Kotwica KA, Ferre CL, Michel GF. Relation of stable
hand-use preferences to the development of skill for
managing multiple objects from 7 to 13 months of
age. Dev Psychobiol 2008, 50:519–529.
139. Yonas A, Hartman B. Perceiving the affordance of
contact in four- and five-month-old infants. Child
Dev 1993, 64:298–308.
140. Chen Z, Siegler RS. Across the great divide: bridging
the gap between understanding of toddlers’ and older
children’s thinking. Monogr Soc Res Child Dev
2000, 65.
141. van Leeuwen L, Smitsman A, van Leeuwen C. Affor-
dances, perceptual complexity, and the development
of tool use. J Exp Psychol Hum Percept Perform
1994, 20:174–191.
142. Rat-Fischer L, O’Regan JK, Fagard J. The emergence
of tool use during the second year of life. J Exp Child
Psychol 2012, 113:440–446.
143. McCarty ME, Clifton RK, Collard RR. Problem sol-
ving in infancy: the emergence of an action plan. Dev
Psychol 1999, 35:1091–1101.
144. Greer T, Lockman JJ. Using writing instruments:
invariances in young children and adults. Child Dev
1998, 69:888–902.
145. Claxton LJ, McCarty ME, Keen R. Self-directed
action affects planning in tool-use tasks with toddlers.
Infant Behav Dev 2009, 32:230–233.
146. McCarty ME, Clifton RK, Collard RR. The begin-
nings of tool use by infants and toddlers. Infancy
2001, 2:233–256.
147. Steenbergen B, van der Kamp J, Smitsman A,
Carson RG. Spoon-handling in two- to four-year-old
children. Ecol Psychol 1997, 9:113–129.
148. Keen R, Lee MH, Adolph KE. Planning an action: a
developmental progression in tool use. Ecol Psychol
2014, 26:96–108.
149. Miller JL, Sonies BC, Macedonia C. Emergence of
oropharyngeal, laryngeal and swallowing activity in
the developing fetal upper aerodigestive tract: an
ultrasound evaluation. Early Hum Dev 2003,
71:61–87.
150. Wilson EM, Green JR, Yunusova YY, Moore CA.
Task specificity in early oral motor development.
Semin Speech Lang 2008, 29:257–266.
151. Burton P, Deng J, McDonald D, Fewtrell MS. Real-
time 3D ultrasound imaging of infant tongue move-
ments during breastfeeding. Early Hum Dev 2013,
89:635–641.
© 2016 Wiley Periodicals, Inc.
The development of motor behavior
152. Geddes DT, Sakalidis VS, Hepworth AR,
McClellan HL, Kent JC, Lai CT, Hartmann PE. Ton-
gue movement and intra-oral vacuum of term infants
during breastfeeding and feeding from an experimen-
tal teat that released milk under vacuum only. Early
Hum Dev 2012, 88:443–449.
153. Barlow SM. Oral and respiratory control for preterm
feeding. Curr Opin Otolaryngol Head Neck Surg
2009, 17:179–186.
154. van der Meer ALH, Holden G, van der Weel FR.
Coordination of sucking, swallowing, and breathing
in healthy newborns. J Pediatr Neonatol 2005,
2:69–72.
155. Fucile S, MacFarland DH, Gisel EG, Lau C. Oral and
nonoral sensorimotor interventions facilitate suck-
swallow-respiration functions and their coordination
in preterm infants. Early Hum Dev 2012,
88:345–350.
156. Sheppard JJ, Mysak ED. Ontogeny of infantile oral
reflexes and emerging chewing. Child Dev 1984,
55:831–843.
157. Wilson EM, Green JR. The development of jaw
motion for mastication. Early Hum Dev 2009,
85:303–311.
158. Reissland N, Francis B, Mason J. Can healthy fetuses
show facial expressions of “pain” or “distress”?
PLoS One 2013, 8:e65530.
159. Reissland N, Francis B, Mason J, Lincoln K. Do
facial expressions develop before birth? PLoS One
2011, 6:e24081.
160. Loos HM, Doucet S, Roussignan R, Hartmann C,
Durand K, Dittrich R, Sagot P, Buettner A, Schaal B.
Responsiveness of human neonates to the odor of 5a-
Androst-16-en-3-one: a behavioral paradox? Chem
Senses 2014, 39:693–703.
161. Messinger DS. Positive and negative: infant facial
expressions and emotions. Curr Dir Psychol Sci
2002, 11:1–6.
162. Messinger DS. Smiling. In: Haith MM, Benson JB,
eds. Encyclopedia of Infant and Early Childhood
Development, vol. 3. Oxford: Elsevier; 2008,
186–198.
163. Messinger DS. Smiling. In: Salkind NJ,
ed. Encyclopedia of Human Development, vol. 3.
Thousand Oakes, CA: Sage; 2006, 1183–1185.
164. Oster H. Emotion in the infant’s face: insights from
the study of infants with facial anomalies. Ann N Y
Acad Sci 2003, 1000:197–204.
165. Green JR, Nip ISB. Some organization principles in
early speech development. In: Maaseen B, van
Lieshout PHHM, eds. Speech Motor Control: New
Development in Basic and Applied Research.
New York: Oxford University Press; 2010, 171–188.
166. Green JR, Moore CA, Higashikawa M, Steeve RW.
The physiologic development of speech motor
 Primer
control: lip and jaw coordination. J Speech Lang
Hear Res 2000, 43:239–255.
167. Gibson JJ. The Ecological Approach to Visual Per-
ception. Boston, MA: Houghton Mifflin Com-
pany; 1979.
168. Fausey CM, Jayaraman S, Smith LB. From faces to
hands: changing visual input in the first two years.
Cognition 2016, 152:101–107.
169. Daniel BM, Lee DN. Development of looking with
head and eyes. J Exp Child Psychol 1990,
50:200–216.
170. Jayaraman W, Fausey CM, Smith LB. The faces in
infant-perspective scenes change over the first year of
life. PloS One 2015, 10:e0123780.
171. Franchak JM, Kretch KS, Soska KC, Adolph KE.
Head-mounted eye tracking: a new method to describe
infant looking. Child Dev 2011, 82:1738–1750.
172. Smith LB, Yu C, Pereira AF. Not your mother’s view:
the dynamics of toddler visual experience. Dev Sci
2011, 14:9–17.
173. Yu C, Smith LB. Multiple sensory-motor pathways lead
to coordinated visual attention. Cognit Sci. In press.
doi: 10.1111/cogs.12366.
© 2016 Wiley Periodicals, Inc.
wires.wiley.com/cogsci
174. von Hofsten C, Rosander K. Development of smooth
pursuit tracking in young infants. Vision Res 1997,
37:1799–1810.
175. Rosander K, von Hofsten C. Development of gaze
tracking of small and large objects. Exp Brain Res
2002, 146:257–264.
176. Rosander K, von Hofsten C. Infants’ emerging ability
to represent occluded object motion. Cognition 2004,
91:1–22.
177. Johnson SP, Amso D, Slemmer JA. Development of
object concepts in infancy: evidence for early learning
in an eye tracking paradigm. Proc Natl Acad Sci
2003, 100:10568–10573.
178. Bronson GW. Infant’s transitions toward adult-like
scanning. Child Dev 1994, 65:1243–1261.
179. Haith MM, Hazan C, Goodman GS. Expectations
and anticipation of dynamic visual events by 3.5-
month-old babies. Child Dev 1988, 59:467–479.
180. Franchak JM, Adolph KE. Visually guided naviga-
tion: head-mounted eye-tracking of natural locomo-
tion in children and adults. Vision Res 2010,
50:2766–2774.